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A New Capybara from the Late Miocene of San Juan Province, Argentina, and Its Phylogenetic Implications

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A New Capybara from the Late Miocene of San Juan Province, Argentina, and Its Phylogenetic Implications A new capybara from the late Miocene of San Juan Province, Argentina, and its phylogenetic implications ESPERANZA CERDEÑO, MARÍA E. PÉREZ, CECILIA M. DESCHAMPS, and VÍCTOR H. CONTRERAS Cerdeño, E., Pérez, M.E., Deschamps, C.M., and Contreras, V.H. 2019. A new capybara from the late Miocene of San Juan Province, Argentina, and its phylogenetic implications. Acta Palaeontologica Polonica 64 (1): 199–212. A new hydrochoerine rodent, Cardiatherium calingastaense sp. nov. (Caviidae), is described based on the specimen INGEO-PV 87. It was recovered from the late Miocene Las Flores Formation, cropping out at the Puchuzum locality, San Juan Province, Argentina. The new species is based on a particular combination of characters, among which the following can be highlighted: p4 with internal fissures equally deep as in Cardiatherium paranense, secondary external fissure as in Cardiatherium patagonicum, and lacking the fifth internal fissure and supernumerary internal fissure, as in C. paranense; m3 with a conspicuous labial column in the posterior ramus of the second prism; very deep primary and secondary external fissures in upper cheek teeth, the former producing a labial strong step-shaped profile in M2; sagittal crest on the parietals; bullae small in ventral view; scars of the origin of the masseter medialis muscle with an anterior projection up to the level of the incisive foramen and the maxilla-premaxilla suture. The phylogenetic analysis supports the taxonomic proposal of creating a new species of Cardiatherium and shows C. calingastaense sp. nov. as the sister group of the other species of the genus. The lineage leading to the clade Cardiatherium + largest capybaras would have originated at least during the Chasicoan SALMA (early late Miocene). Cardiatherium calingastaense sp. nov. adds to the previous record of Cardiatherium chasicoense and Cardiatherium paranense in the late Miocene of San Juan and Mendoza provinces, respectively, thus increasing the diversity of capybaras in central-west Argentina. Key words: Mammalia, Rodentia, Caviidae, Cardiatherium, systematics, phylogeny, Miocene, Argentina. Esperanza Cerdeño [[email protected]], Instituto Argentino de Nivología, Glaciología y Ciencias Ambi- entales (IANIGLA), Centro Científico Tecnológico, CONICET Mendoza, Av. Ruiz Leal s/n, 5500, Mendoza, Argentina. María E. Pérez [[email protected]], Museo Paleontológico Egidio Feruglio, Av. Fontana 140, U9100GYO, Trelew, Argentina. Cecilia M. Deschamps [[email protected]], Comisión de Investigaciones Científicas de la provincia de Buenos Aires, Argentina; División Paleontología Vertebrados, Unidades de Investigación Anexo Museo de La Plata, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, 122 y 60, 1900, La Plata, Argentina. Víctor H. Contreras [[email protected]], Instituto de Geología Dr. Emiliano P. Aparicio (INGEO) y Depar- tamento Geología, FCEFN, Universidad Nacional de San Juan, Ignacio de la Roza y Meglioli, 5400 Rivadavia, San Juan, Argentina. Received 5 September 2018, accepted 19 October 2018, available online 26 November 2018. Copyright © 2018 E. Cerdeño et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License (for details please see http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. and the Huayquerian South American Land Mammal Ages Introduction (SALMAs; Contreras et al. 1995; Peralta et al. 1995; Contreras The Neogene continental outcrops in the San Juan Province, 1996; Cerdeño and Contreras 2000; Ciancio et al. 2006; León central-west Argentina (Fig. 1), encompass two localities and Bracco 2008; Bracco et al. 2011; Contreras and Baraldo that have yielded important late Miocene vertebrate remains. 2011; Suvires and Contreras 2011). These localities are Puchuzum (Calingasta Valley), whose Concerning the fossils from Puchuzum, different lev- faunal assemblage is assigned to the Huayquerian SALMA els of the section (distinct architectural elements) have (late Tortonian–Messinian; see Deschamps et al. 2013, for provided remains of molluscs, ostracods, insects, fishes, further details in bio- and chronostratigraphic equivalences), and palynomorphs (found within lacustrine sediments), and Loma de Las Tapias, where two different levels are on the one hand, and mammals and birds (derived from recognized as corresponding to the Chasicoan (Tortonian) flood plain sediments), together with footprint tracks, on Acta Palaeontol. Pol. 64 (1): 199–212, 2019 https://doi.org/10.4202/app.00544.2018 200 ACTA PALAEONTOLOGICA POLONICA 64 (1), 2019 A B W6933’ N 25 Casta o viejo River 35 ARGENTINA Candelaria Creek S3106’36’’ Puchuzum 45 500 km 55 1km 80 70 60 Fig. 1. Geographical map showing position of studied area within San Juan Province, central-west Argentina (A) and location of outcrops of the Las Flores Formation (striped area) near the village of Puchuzum (B). The asterisks indicate the fossil locality. the other hand (Contreras 1996; Contreras and Peralta 1998; 1883a, b; Rovereto 1914; Kraglievich 1930; Rusconi 1933, Prámparo et al. 1998; Contreras and Cerdeño 2010; Bracco 1935, 1944; Pascual and Bondesio 1963; Francis and Mones et al. 2011). Among mammal remains, only the specimens 1965a, b; Bondesio 1985a, b; Mones 1991; among others), of a Hegetotheriidae (Notoungulata) were studied in de- but revisions such as those by Mones (1984) and, more tail (Cerdeño and Contreras 2000). However, the faunal recently, Vucetich et al. (2005, 2012, 2014a, b, 2015) and assemblage also includes xenarthrans, litopterns and ro- Deschamps et al. (2007, 2009) have evidenced a great taxo- dents (Contreras and Cerdeño 2010; Bracco et al. 2011). nomic overestimation due to ontogenetic variation that had The hegetotheriid Hemihegetotherium achataleptum indi- not been previously evaluated; consequently, these authors cated a Huayquerian SALMA for this fauna (Cerdeño and drastically reduced the known diversity of hydrochoerines, Contreras 2000), but no absolute age has been obtained for especially for the late Miocene–Pliocene interval. the unit. Herein we present the study of a well-preserved Within the late Miocene, the oldest species is Cardia- skull and mandible of a capybara, associated with some therium chasicoense Pascual and Bondesio, 1968 from the fragments of postcranial bones. Chasicoan SALMA, which is known from the type locality Capybaras are the largest living rodents, represented by Arroyo Chasicó (Buenos Aires Province) as well as from the single genus Hydrochoerus, reaching a body length of the lower levels of the Loma de Las Tapias Formation in 100–130 cm, and adult males weighting up to 91 kg (Mones San Juan Province (Contreras 2000; Contreras et al. 2001; and Ojasti 1986). They have semiaquatic habits, and live Deschamps et al. 2007, 2009; Contreras and Baraldo 2011). in groups usually ranging in size from two to 30 individ- In turn, the Huayquerian SALMA localities with hydro- uals (Dunnum 2015). They have characteristic euhypsod- choerines are more abundant and the material was re-eval- ont, multilaminated cheek teeth (Novak 1991; Deschamps et uated by Vucetich et al. (2005). These authors described al. 2007). The phylogenetic relationships of capybaras with Cardiatherium patagonicum as the southernmost record, other cavioid rodents and the status of hydrochoerids have from the Puerto Madryn Formation (Chubut Province). been revised during the last years (Rowe and Honeycutt At the same time, they proposed that all material from 2002; Vucetich and Pérez 2011; Upham and Patterson 2015; the Ituzaingó Formation (Entre Ríos Province, north- Pérez et al. 2018, and references therein). After these au- east Argentina) would correspond to only one species, C. thors, capybaras would be within the Family Caviidae as a paranense, which was later confirmed by Vucetich et al. subfamily, Hydrochoerinae, together with the extinct car- (2014b). These authors also recognized C. paranense in diomyines and the small Kerodon (rock-dwelling mocos), Tupungato, Mendoza Province (central-west Argentina), the other living genus of the subfamily (Rowe and Honeycutt from levels of the Río de Los Pozos Formation, referred to 2002; Pérez and Pol 2012; Pérez et al. 2018). the late Miocene (Yrigoyen 1993). Another species assigned The fossil record of capybaras (that is, hydrochoerines to this genus, Cardiatherium orientalis Francis and Mones, excluding ‘cardiomyines’) goes back to the late Miocene, and 1965b, was recorded in the late Miocene of Uruguay, and many taxa were described since the 19th century (Ameghino tentatively recognized later in the Cerro Azul Formation, La CERDEÑO ET AL.—A NEW MIOCENE CAPYBARA FROM ARGENTINA 201 Pampa Province, Argentina, as C. aff. orientalis (Vucetich Nomenclatural acts.—This published work and the nomen- et al. 2014b). The genus Cardiatherium is also known in the clatural acts it contains, have been registered in ZooBank: late Miocene of Brazil, Peru and Venezuela (Deschamps et urn:lsid:zoobank.org:pub:2A1D2DDE-7100-4BC7-B0B1- al. 2013 and literature therein). 8862CC1A064E. The present contribution focuses on the description of a new species of Cardiatherium based on well-preserved cranial remains, previously mentioned as Cardiatherium sp. Material and methods (Contreras and Cerdeño 2010), found in the province of San Juan. On the basis of a cladistic analysis, the phylogenetic The studied material,
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